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Biogeography and Evolution of the
Desert Stink Beetles
(Coleoptera: Tenebrionidae: Amphidorini)
M. Andrew Johnston
AmphidoriniLeConte, 1862*
• New World tribe
• 7 Genera
– 17 subgenera of Eleodes Eschscholtz
• 256 valid species/subspecies
– 213 in Eleodes, with 163 additional
synonyms
= Eleodini Blaisdell, 1909
Eleodes spinipes macrura Champion, 1892
AmphidoriniLeConte, 1862
• Last comprehensive revision was
Blaisdell, 1909
• Treated 115 species/subspecies
from the U.S. and Canada
Eleodes snowi Blaisdell, 1909
AmphidoriniLeConte, 1862
• All species are flightless
• Known for defensive “head-
standing” posture
Eleodes hispilabris (Say, 1824)
Several sympatric species found in Arizona
E. madrensis Johnston
E. subnitens LeConte E. longicollis LeConteE. carbonaria Say
E. anthracina Blaisdell
Eleodes Eschscholtz, 1829
Müllerian mimicry
Collected at same time
(Heber, AZ 2013)
Smith et al. 2014, fig. 2
Morphological
convergence
• Sculpturing
• Dorsal silhouette
Batesian Mimicry
• 2 are Stenomorpha
• Lack defensive
glands
• Mimic
headstanding
behavior
• ~180 my divergence
Collected at same time
(Heber, AZ 2013)
Smith et al. 2014, fig. 2
Biodiversity data and darkling beetles
Eleodes Eschscholtz, 1829
213 Valid species-group taxa
412 Available species-group names
Eleodes species in Arizona
34 verified
4 doubtful
38
38
8,010 Eleodes specimens
from Arizona
38
208
Darwin Core ‘scientificName’
208 unique strings (“names”)
38
208
Darwin Core ‘scientificName’
Eleodes carbonaria chihuahuaensis
Champion, 1884
Previously valid combinations
eleodes carbonarius chihuahuaensis
eleodes lineatus
eleodes ampla
eleodes nitidus
eleodes lineata
eleodes amplus
eleodes carbonarius amplus
eleodes amplus dolosus
eleodes ampla dolosa
eleodes nitida
Taxonomic Issues I – Synonymy
38
208
Darwin Core ‘scientificName’
Eleodes carbonaria chihuahuaensis
Champion, 1884
Subgeneric names included
eleodes (melaneleodes) amplus
eleodes melaneleodes lineata
eleodes melaneleodes amplus
Taxonomic Issues II – Dirty Data
38
208
14
‘Accepted’ species from
backbone taxonomy
8 verified
6 doubtful
38
208
14
Manually interpreted using
current literature
32 verified
14 doubtful
51
38
208
14
Good data are available
But we need a good taxonomy
to realize their potential
51
2315 Eleodes records
74 species/subspecies
2315 Eleodes records
74 species/subspecies
Default view of data
705 specimens
“ID’d” to Eleodes
29 taxon names
recognized
Verbatim view of record, showing issue flags
Failure across (at least) Coleoptera
iPhylo Blog Post
iPhylo.blogspot.com
~85,000 Australian Coleoptera records
1/3 names are changed, affecting 1/5 records
Graduate students should be empowered
to use and curate biodiversity data
Backbone taxonomies and early-career systematists
Graduate students should be empowered
to use and curate biodiversity data
Backbone taxonomies and early-career systematists
Current backbone taxonomies alienate systematists
Graduate students should be empowered
to use and curate biodiversity data
Backbone taxonomies and early-career systematists
Need to develop new, community-empowering designs
Current backbone taxonomies alienate systematists
Tenebrionidae of the Algodones Dunes
Tenebrionidae of the Algodones Dunes
The Coleopterous fauna of selected
California sand dunes
Andrews et al. 1979
23 species reported
Updated ‘expert-asserted’ checklist
54 species reported
Updated ‘expert-asserted’ checklist
54 species reported
Explicit
taxon concept
Updated ‘expert-asserted’ checklist
54 species reported
Explicit
taxon concept
Cited but vague
data sources
Checklist generated from digitized records
Checklist generated from digitized records
Expert-asserted
54 verified
Checklist generated from digitized records
52 taxa listed
31 verified
18 ‘synonyms’
3 ‘mis-IDs’
Checklist generated from digitized records
45 taxa listed
25 verified
18 ‘synonyms’
2 ‘mis-IDs’
Checklist generated from digitized records
17 taxa listed
15 verified
2 ‘synonyms’
Checklist generated from digitized records
Currently digitized data are useful, though not complete
Checklist generated from digitized records
Currently digitized data are useful, though not complete
Faunal and revisionary studies should
explicitly reference available records
Checklist generated from digitized records
Currently digitized data are useful, though not complete
Faunal and revisionary studies should
explicitly reference available records
Taxonomic expertise is still very much required
Back to the desert stink beetles…
Neatus_n_sp_TB15296
Lariversius_tibialis_BigDune_E79
Promus_knullorum_E24
Nycterinus_sp3_ElFaro_TB15413
Eleodes_sp6_TB15084
Eleodes_dentipes_TB15297
Litheleodes_arcuatus_E140
Caverneleodes_n_sp_1_Tonto_E25
Eleodes_eschscholtzi_E21
Pimeliotus_lugens_TB14731
Melaneleodes_wenzeli_E15
Eleodes_debilis_TB15710
Carchares_macer_E149
Eleodes_unknownB_E171
Eleodes_rileyi_TB15994
Trogloderus_Olancha_E81
Ardeleodes_tibialis_E145
Blapylis_sp_TulareCo_E142
Promus_hogei_E40
Cratidus_osculans_E36
Eleodes_barbatus_TB15989
Omegeleodes_puebla_sp_E167
Discogenia_scabriculus_E67
Eleodes_littoralis_Eschscholtz_1831_Amphidora_TB15447
Rhinandrus_sp_TB15991
Nycterinus_sp2_Cartagena_TB15412
Blaps_mucronota_TB15113
Eleodes_sp3_TB15081
Eleodes_labialis_TB15486
Rhinandrus_sp_TB15987
Eleodes_quadricollis_TB15293
Ammodonus_granosus_E129
Eleodes_acuticaudus_E34
Steneleodes_distinctus_E154
Conibius_sp_E139
Steneleodes_sallaei_E169
Eleodes_clavicocnis_E83
Eleodes_armatus_E13
Eleodes_impolitus_TB15074
Eleodes_tenuipes_TB15993
Zophobas_morio_TB15298
Eleodes_distinctus_TB15079
Embaphion_elongatum_TB15300
Trogloderus_costatus_E60
Rhinandrus_helopioides_E65
Eleodes_sp9_TB15114
Eleodes_barbatus_TB15992
Nycterinus_sp4_TB15448
Heteropromus_veterator_E74
Anomalipus_sp_TB14734
Trichoton_sp1_Nacunan_TB13471
Steneleodes_pr_coarctatus_E172
Embaphion_muricatum_E51
Eleodes_Pseudeleodes_sp_KKDNA0392
Metablapylis_delicatus_E141
Eleodes_puebla_sp_E162
Embaphion_contusum_E46
Eleodes_80_E37
Eleodes_grandicollis_TB13402
Steneleodes_laevigatus_E174
Promus_oaxaca_sp_E165
Melaneleodes_carbonarius_E14
Steneleodes_mutilatus_E9
Melaneleodes_carb_chihuahuaensis_E136
Blapylis_nigropilosus_E90
Argoporis_sp2_Texas_TB15454
Melaneleodes_tricostatus_E68
Steneleodes_hepburni_E8
Eleodes_obs_sulcipennis_E11
Omegeleodes_oaxaca_sp_E159
Zophobas_sp_2_Chiapas_TB15985
Lariversius_kelso_TB13435
Nycterinus_sp2_E75
Scotobius_sp2_TB15190
Eleodes_sp4_TB15082
Promus_insularis_E54
Tricheleodes_hirsutus_E164
Embaphion_depressum_E73
Metablapylis_nigrinus_E38
Steneleodes_gravidus_E30
Eleodes_Promus_insularis_TB15981
Centronopus_suppressus_E71
Epantius_obscurus_TB15187
Eleodes_loretensis_TB15485
Trogloderus_NMSanJuan_E91
Omegeleodes_debilis_E64
Eleodes_Steneleodes_sp_TB15477
Embaphion_glabrum_E50
Eleodes_Eleodes_loretensis_TB15982
Eleodes_scapularis_E158
Zophobas_sp_1_Ecuador_TB15984
Leptynoderes_sp1_TB15192
Steneleodes_impolitus_E166
Eleodes_Eleodes_armatus_TB13139
Neobaphion_papula_E157
Eleodes_Eleodes_grandicollis_TB15986
Eleodes_spinipes_macrurus_TB15473
POI7_E156
Eleodes_nr_dentipes_TB15474
Eleodes_tibialis_TB15112
Argoporis_cribrata_E56
Eleodes_Melaneleodes_debilis_TB13393
Notibius_puberulus_E138
Metablapylis_n_sp_teelsmarsh_E175
Promus_alutaceus_E49
Steneleodes_angustus_E168
Eleodes_peropacus_TB15476
Eleodes_pilosus_Horn_1870_Eleodes_TB15449
Eleodes_sp2_TB15080
Blaps_sp1_TB17179
Eleodes_dissimilis_TB15478
Eleodes_sp_Oaxaca_E146
Eleodes_sp8_TB15077
Promus_calcaratus_E163
Cymatothes_opacus_E72
Eleodes_marginatus_TB15890
Eleodes_impolitus_E53
Cerenopus_concolor_TB15484
Eleodes_longipilosus_TB15304
Gonopus_sp1_TB14732
Melaneleodes_carb_obsoletus_E42
Caverneleodes_wynnei_E26
Lariversius_tibialis_E70
Embaphion_sp1_Carlsbad_TB15451
Cheirodes_sp_Nabibia_E148
Emmallodera_sp1_TB15191
Blapylis_tribulus_E61
Melaneleodes_carb_omissus_E12
Eleodes_spinipes_ventriculosus_E18
Litheleodes_extr_TXElPaso_E52
Promus_fusiformis_E44
Eleodes_littoralis_Eschscholtz_1831_Amphidora_TB15446
Eleodes_sp5_TB15083
Eleodes_acutus_TB17182
Eleodes_sp_nr_pilosus_TB15305
Eleodes_sponsus_E39
Eleodes_spinipes_spinipes_E155
Eleodes_longicollis_TB15295
Lariversius_eureka_TB15302
Apsena_sp1_TB15455
Steneleodes_ponderosus_E22
Apsena_sp2_TB15456
Eleodes_sallaei_var_2_TB15075
Litheleodes_extricatus_AZNav_E62
Eleodes_suturalis_E19
Eleodes_mexicanus_TB15483
Steneleodes_innocens_E41
Eleodes_Steneleodes_TB15303
Steneleodes_Hidalgo_sp_E160
Ele_sp1_TB13388
Eleodes_gracilis_E28
Eleodes_tenuipes_E17
Eleodes_aristatus_TB13389
Eleodes_Melaneleodes_debilis_TB15988
Blapylis_snowi_E31
Eleodes_sp_Tlaxcala_E132
Eulabs_bicarinata_TB15299
Eleodes_sp7_TB15085
Eleodes_rossi_E35
Promus_madrensis_E128
Chaseleodes_connatus_E58
Steneleodes_longicollis_E16
Promus_subnitens_E10
Promus_goryi_E27
Ele_sp81_1_TB13394
Eleodes_ponderosus_TB15078
Nycterinus_sp1_Tiltil_TB15411
Blaps_sp3_TB17181
Eleodes_caudiferus_E29
Eleodes_esch_eschscholtzi_E23
Neobaphion_planipennis_E43
Eleodes_hirsutus_TB15479
Steneleodes_ruidus_E33
Promus_striolatus_E80
Blaps_sp2_TB17180
Eleodes_acuticaudus_LeConte_TB15444
Eleodes_spinipes_spinipes_TB15076
Litheleodes_extr_TXWard_E48
Melaneleodes_anthracinus_E173
Lariversius_tibialis2_E57
Eleodes_opacus_TB17183
Steneleodes_giganteus_E20
Blapylis_tenebrosus_E47
Trogloderus_costatus_costatu_E176
Omegeleodes_oaxaca_sp2_E161
Eleodes_sp_TB15481
Metablapylis_dissimilis_E7
Eleodes_giganteus_TB15475
Eleodes_nr_distinctus_E144
Unk_larvae_Tiltil_TB15420
Apsena_rufipes_TB15409
Scotobius_sp_TB15115
Rhinandrus_sp1_Peru_TB15445
Litheleodes_extricatus_AZGraham_E63
Eleodes_Eleodes_loretensis_TB15983
Tricheleodes_pilosus_E45
Tricheleodes_barbatus_E59
Embaphion_NMSmall_TB17171
Promus_spinolae_E32
Caverneleodes_microps_E76
Gonopus_sp1_E150
Steneleodes_UnknownA_desierto_liones_E170
Eleodes_loretensis_TB15674
Litheleodes_extr_NavajoNM_E55
Blapstinus_fortis_E69
Nycterinus_sp3_E151
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100
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46
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60
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9
100
63
42
20
13
29
100
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100
97
100 24
100
74
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52
62
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44
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52
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32
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8
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20
Phylogenetic analysis of Amphidorini
Smith et al. in prep, ML tree
Maximum Likelihood
Tree
(186 terminals, 7 loci, RAxML)
Current Generic Classification
Phylogenetic analysis of Amphidorini
Current Generic Classification
Phylogenetic analysis of Amphidorini
South American Nycterinus Eschscholtz
does not belong in Amphidorini
‘True Amphidorini’ are only found in
North America
Nycterinus rugiceps Curtis, 1845
Phylogenetic analysis of Amphidorini
Paraproct
Valvifer
(Coxite I)
Gonostyle
Coxite IV
Coxite III
Coxite II
Nycterinus rugiceps Curtis, 1845 Eleodes armata LeConte, 1851
Current Generic Classification
Phylogenetic analysis of Amphidorini
Remaining genera are derived within the
large genus Eleodes Eschscholtz
Current Generic Classification
Phylogenetic analysis of Amphidorini
Eleodes hepburni Champion, 1884 Eleodes carbonaria Say, 1823Trogloderus n.sp.
Consensus tree of 500 ML replicates
7-Gene Phylogeny Backbone
is Poorly Supported
7-way Polytomy 5-way Polytomy
Phylogenetic analysis of Amphidorini
Supports our 7-gene
topology
Currently analyzing
29 transcriptomes
from all major
lineages
Preliminary 343-loci analysis
Phylogenetic analysis of Amphidorini
Revised Generic Classification
Phylogenetic analysis of Amphidorini
14 Genera (only 1 without a name)
12 Eleodes subgenera (4 unnamed)
Comparative Morphology
Many molecular clades are
finding morphological support
Female ovipositor characters
can diagnose subgenera
Female Ovipositors of 3 different subgenera,
A,B: E. tribula, C: E. barbata, D: E. hirsuta
Historical Biogeography
From Zhang et al.
2016
Historical Biogeography
From Zhang et al.
2016
1 – Species distributional data
Historical Biogeography
From Zhang et al.
2016
1 – Species distributional data
2 – Phylogeny
Historical Biogeography
From Zhang et al.
2016
1 – Species distributional data
2 – Phylogeny 3 – Areas of Endemism
CORRELATION*
Analyzes digitized herbarium data from SEINet
*Landrum, L.R. and D. Lafferty. 2015. Taxon 64(5) 998-1016
http://dx.doi.org/10.12705/645.9
CORRELATION*
*Landrum, L.R. and D. Lafferty. 2015. Taxon 64(5) 998-1016
http://dx.doi.org/10.12705/645.9
Computes how frequently each species is found
near each other given species
CORRELATION*
*Landrum, L.R. and D. Lafferty. 2015. Taxon 64(5) 998-1016
http://dx.doi.org/10.12705/645.9
Clusters species into co-occurring groups
CORRELATION
*Brown, D.E. (ed.) 1994. Biotic communities: Southwestern United States and northwestern
Mexico. Salt Lake City: University of Utah Press.
Arizona Flora Test Case:
• Examined 81 ‘high
profile’ species
• Recovered groupings
highly congruent with
published biotic
communities*
CORRELATION
Analyze input set of
lat/long coordinates:
• Correlate an animal
species with SEINet
plants
• Place that animal in a
plant-driven biotic
community
Using CORRELATION, we can
ask questions about ecological
communities
And answer them based on
specimen occurrence data
Where can we find a certain species?
“You can find it in
oak-juniper forests in
southern Arizona”
Eleodes madrensis Johnston, 2015
Where can we find a certain species?
Eleodes madrensis Johnston, 2015
5 highest correlated plant
species
“You can find it in
oak-juniper forests in
southern Arizona”
Differentiate habitats of similar species
Eleodes arcuata Casey, 1884Eleodes debilis Horn, 1870
Differentiate habitats of similar species
Eleodes arcuata Casey, 1884Eleodes debilis Horn, 1870
Differentiate habitats of similar species
Eleodes arcuata Casey, 1884Eleodes debilis Horn, 1870
Only share a single species of top 10
Differentiate habitats of similar species
Eleodes arcuata Casey, 1884
Eleodes arcuata is
correlated with oak-
juniper mid-elevation
transition forest
Differentiate habitats of similar species
Eleodes debilis Horn, 1870
Eleodes debilis is
correlated with riparian
broadleaf plants from a
similar elevation
CORRELATION transforms our approach to
understanding distribution
Ecological Community
“Authoritative observation”
Repeatable, specimen-based
correlation
Ecological Community
“Authoritative observation”
Repeatable, specimen-based
correlation
Spatial Distribution
Geographic polygons
Specimen-driven, co-occurring
species communities
CORRELATION transforms our approach to
understanding distribution
Can we infer ancestor-descendant
plant-community shifts?
Historical plant-community inference
• 21 Eleodes species from Arizona were selected
– Records with high geographic uncertainty or missing
locality data were removed
– Records were sorted to taxa by ‘ScientificName’ based on
nomenclatural synonymies
Historical plant-community inference
• 21 Eleodes species from Arizona were selected
• Associated beetles with plant communities using
CORRELATION
– Beetle species associated with 6 different plant
communities
– 18 species were correlated to a single plant community, 3
were equally correlated to 2 communities
Historical plant-community inference
• 21 Eleodes species from Arizona were selected
• Associated beetles with plant communities using
CORRELATION
• ‘Biogeographic’ reconstruction in BioGeoBEARS
– Inferred historical plant-community lineage associations
– Assumed current plant communities existed throughout
Eleodes evolutionary history
Historical plant-community inference
Colors represent six
different biotic
communities
Phylogeny based on a 7-gene unpublished
dataset, Smith et al. in prep
Historical plant-community inference
Two independent
radiations into the
Sonoran Desert
community
Phylogeny based on a 7-gene unpublished
dataset, Smith et al. in prep
Historical plant-community inference
‘Melaneleodes Clade’
ancestors associated
with Pine-Juniper
community of the
Colorado Plateau
Phylogeny based on a 7-gene unpublished
dataset, Smith et al. in prep
Historical plant-community inference
Ancestral lineage
associated with higher
elevation oak-juniper
and ponderosa pine
communities
Phylogeny based on a 7-gene unpublished
dataset, Smith et al. in prep
• Novel analysis for inferring historical habitat
transitions
• Entirely based on vouchered specimen data
• Analyses are reproducible and easily re-done when
new data are made available
Historical plant-community inference
Trogloderus LeConte, 1879
• Endemic to sand dunes
and sandy habitats in
western United States
• Currently comprised of 1
species with 4 subspecies
Trogloderus ‘major’ n.sp.
Last revised by La Rivers in
1946
• Invoked orthogenetic evolution
Last revised by La Rivers in
1946
• Invoked orthogenetic evolution
• Hypothesized Trogloderus to be ancient
Last revised by La Rivers in
1946
• Invoked orthogenetic evolution
• Hypothesized Trogloderus to be ancient
• Hypothesized origins from the Great Basin
Phylogenetic revision of Trogloderus
36 OTUs, 6 loci
analyzed in BEAST
• 10 total species
• 4 described, 6 new
• Trogloderus perhaps
~5.5 my old
Trogloderus root
Distributed through the
Intermountain region
But …
How do we divide up areas
of endemism?
Trogloderus Biogeography
Classical, simplistic view
U.S. EPA Ecoregions
Van Dam and Matzke 2016
What about classifying sand dune regions?
Trogloderus as a biogeographic model
• Moderate habitat specificity
• Sandy, well-drained soils
• Apparently wide temperature tolerance
• Low to moderate dispersal
• Flightless
• Not restricted to single localities
• Species show strong geographic signal
Great Basin
T. costatus LeConte
Great Basin
T. nevadus La Rivers
Great Basin
T. costatus LeConte
Great Basin
T. nevadus La Rivers
Prehistoric Lake Subregions
• Lake Lahontan (W)
• Lake Bonneville (E)
Great Basin – Mohave Transition
T. ‘kandai’ n. sp.
T. ‘arcanus’ n. sp.
T. ‘major’ n. sp.
Great Basin – Mohave Transition
T. ‘kandai’ n. sp.
T. ‘arcanus’ n. sp.
T. ‘major’ n. sp.
“Great Basin”
“Mohave Desert”
Lahontan Trough
Proposed by Reveal in 1979
• Low-elevation desert
corridor
• Geologically distinct from
Lake Lahontan
Lahontan Trough
Eusattus muricatus LeConte
• Genetically distinct population1
• First to recognize Lahontan Trough
as a genetic barrier
1Britten & Rust 1996. Conservation Biology
Lahontan Trough
Eusattus muricatus LeConte
• Genetically distinct population1
• First to recognize Lahontan Trough
as a genetic barrier
E. muricatus species group
• Exhibit speciation consistent with
the Lahontan Trough2
1Britten & Rust 1996. Conservation Biology
2Doyen 1984. Occ. Papers of Cal. Acad. Sci.
Owens Lake Basin
T. ‘kandai’ n. sp.
T. ‘arcanus’ n. sp.
T. ‘major’ n. sp.
“Great Basin”
“Mohave Desert”
• Southern extent of ‘Great
Basin’ flora
• Not connected to Lahontan
Trough
• T. ‘kandai’ is the first
reported endemic sand
obligate
Originated in Great
Basin, 5-6 mya
Historical Biogeography
Historical Biogeography
• Geographically-linked
speciation events
• About 2-4mya
• Consistent with
separate drainage
systems which didn’t
overlap during the wet
pleistocene
Revisiting La Rivers 70 years later
• Invoked orthogenetic evolution
Revisiting La Rivers 70 years later
• Invoked orthogenetic evolution
– Well-adapted to cryptic living in sand
Revisiting La Rivers 70 years later
• Invoked orthogenetic evolution
– Well-adapted to cryptic living in sand
• Hypothesized Trogloderus to be ancient
Revisiting La Rivers 70 years later
• Invoked orthogenetic evolution
– Well-adapted to cryptic living in sand
• Hypothesized Trogloderus to be ancient
– Likely relatively young, roughly 5–6my old
Revisiting La Rivers 70 years later
• Invoked orthogenetic evolution
– Well-adapted to cryptic living in sand
• Hypothesized Trogloderus to be ancient
– Likely relatively young, roughly 5–6my old
• Hypothesized origins from the Great Basin
Revisiting La Rivers 70 years later
• Invoked orthogenetic evolution
– Well-adapted to cryptic living in sand
• Hypothesized Trogloderus to be ancient
– Likely relatively young, roughly 5–6my old
• Hypothesized origins from the Great Basin
– Probably true; Great Basin younger than he knew
Trogloderus Biogeography Insights
• Intermountain Region has produced complex
speciation during the recent past
• Transition zones between broader ecoregions
could be their own areas of endemism and
speciation
Collaborators
Nico Franz
Sal Anzaldo
Andrew Jansen
Brian Reiley
Bill Warner
Fred Skillman
Kojun Kanda
Aaron Smith
Sangmi Lee
Liz Makings
Chris Wirth
Ryan Lume
Travis Hitchner
Brennan Hayes
David Flemming
Monica Chacon
CMN Visiting Scientist Fellowship
CanaColl
Foundation
NSF ARTS DEB-1258154
Funding
Thank you!

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Biogeography and Evolution of the Desert Stink Beetles

Editor's Notes

  1. Doyen and Somerby 1974
  2. Perhaps the most majestic group of all animals Grammatical gender – nomenclaturally complex group
  3. Header – taxonomic historiy/issues etc Issues I, then II next slide Mention literal text copy
  4. Harvested and available via GBIF
  5. Verbatim records are available, but then you need taxonomic expertise to decode Not issues with data quality, but with backbone taxonomic filtering EARLY CAREER SYSTEMATISTS - FRUSTRATION
  6. Building global initiatives / infrastructure – should be used by and encourage participation from E.C. researchers
  7. ~70km long (up to 10km wide)
  8. Mt genes: COI(JP), COII, 12s NPC genes: H3, wnt, CAD1, 28s
  9. Legend
  10. ADD DIVISION GRAPHICS
  11. ----- Meeting Notes (11/6/14 12:06) ----- Explain orthogenic, or do not use
  12. ----- Meeting Notes (11/6/14 12:06) ----- Explain orthogenic, or do not use
  13. ----- Meeting Notes (11/6/14 12:06) ----- Explain orthogenic, or do not use
  14. ----- Meeting Notes (11/6/14 12:06) ----- Explain orthogenic, or do not use
  15. ----- Meeting Notes (11/6/14 12:06) ----- Explain orthogenic, or do not use
  16. ----- Meeting Notes (11/6/14 12:06) ----- Explain orthogenic, or do not use